2020 05 07 051037v2 Full

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Cast away on Mindoro island: population

2 dynamics of the critically endangered tamaraw
(Bubalus mindorensis) at Mounts Iglit–Baco
4 Natural Park
C. Bonenfant1 , A. Rutschmann2 , J. Burton3 , R. Boyles4 , and

E. Schütz5
1 Université de Lyon, F-69 000, Lyon ; Université Lyon 1 ; CNRS, UMR 5558,
Laboratoire ”Biométrie et Biologie Évolutive”, F-69 622, Villeurbanne, France.
2 School of Biological Sciences, University of Auckland, Auckland, New Zealand
3 IUCN SSC Asian Wild Cattle Specialist Group, Global Wildlife Conservation,
Texas, USA
4 Department of Environment and Natural Resources of the Philippines,
Occidental Mindoro
5 D’Aboville Foundation and Demo Farm Inc, Manila, Philippines
6 Keywords
counts, density-dependence, exponential model, Mindoro dwarf buffalo, protection
8 measures, poaching, population growth.
Correpondence
10 Christophe Bonenfant, christophe.bonenfant@univ-lyon1.fr – ORCID
0000-0002-9924-419X
bioRxiv preprint doi: https://doi.org/10.1101/2020.05.07.051037; this version posted April 1, 2021. The copyright holder for this preprint
C. Bonenfant Tamaraw dynamics on Mindoro
Abstract
2 Endangered species, despite living at low population density, may undergo

density-dependent feed-backs in case of successful recovery or marked reduction
4 in range. When at work, density-dependence dynamics increases extinction risks
and hamper conservation efforts. The largest population of the critically
6 endangered tamaraw (Bubalus mindorensis) lives on a < 3, 000ha in Mounts
Iglit-Baco Natural Park (MIBNP) with very limited expansion possibilities.
8 Tamaraw abundance has been monitored on a yearly basis from animal counts by
Philippine authorities since the year 2000. Consistent with its protected status by
10 law, the MIBNP tamaraw population has been increasing in size at an average
rate of +6% per year, which we found to be relatively low compared to other
12 similar-sized Bovinea species. Population growth was strikingly spatially
structured within MIBNP with a population growth close to +16% in the core
14 area of protection, while a reduction of abundance of −27% was measured at the
periphery of the species range inside MIBNP. Highly concerning is the fact that
16 the annual population growth rate progressively decreased significantly over the
years since 2008, which we interpreted as an evidence of density-dependence.
18 This isolated tamaraw population is currently experiencing a contraction of its
range in MIBNP, likely caused by anthropogenic pressures forcing large
20 herbivores to live at relatively high density in the core zone of the monitoring
where protection is most effective. Our study highlights that beyond the
22 encouraging results of a continuous growth over the last two decades, the
MIBNP tamaraw population remains subject to uncertainty of its long term
24 viability.
2
INTRODUCTION
2 In conservation planning, population abundance is critical and focuses the

attention of field biologists and wildlife management institutions. Low
4 population abundance is considered as one of the foremost criteria used for the
evaluation of conservation statuses (Mace et al. 2008; Neel et al. 2012) . The
6 International Union for Conservation of Nature (IUCN) would for instance
classify any species with very small population sizes as threatened, and so
8 without evidence for past, present or future decline (Mace et al. 2008). Also
most populations of endangered wild species are indeed isolated and fragmented,
10 and hence are of small size. The repeated focus on abundance in conservation
biology has its roots in the fact that population viability strongly depends on the
12 number of animals in a population (Lande et al. 2006). Overall, the ecology of
small populations is particularly relevant to the genetics, evolution and
14 demography of threatened species (Korn 1994).
In terms of populations dynamics, small populations differ in several ways
16 from what is generally reported at higher abundance (Caughley 1994; Mugabo
et al. 2013). When population size is small, the random death of a few
18 individuals may precipitate the extinction of a population, a process referred to
as demographic stochasticity (Shaffer 1981; Lande 1993). Similarly, when
20 mating partners are too few to meet and reproduction fails, demographic
stochasticity generates the so-called Allee effects (Courchamp et al. 1999), that
22 is a decrease of the population growth at low population abundance (i.e.
demographic component, see Stephens et al. (1999)). Yet vulnerable populations
24 can also suffer from classical density-dependent effects whereby population
3
growth rate decreases with animal density (Nicholson 1933), driving the fate and
2 viability of populations at risk of extinction (Beissinger & Westphal 1998; Henle
et al. 2004) . The classical density-dependent effects on population dynamics
4 and demographic rates are often overlooked in conservation because they are a
priori expected to occur mainly at high population abundance. Although
6 counter-intuitive for threatened populations, increasing abundance could,
however, occur incidentally following a range contraction caused by habitat loss,
8 the continuous and long-term reintroduction of individuals for reinforcement, or
following successful protection measures. High abundance can potentially
10 trigger undesirable ecological and demographic consequences for its
conservation.
12 Density-dependence is a pervasive demographic responses among large
mammals (Fowler 1987; Bonenfant et al. 2009). In addition to depressing
14 recruitment rates rapidly, mortality of adults may increase as population density
approaches the carrying capacity (Eberhardt 1977; 2002). Many changes in
16 behaviour and life history traits of individuals account for those observed
changes in demographic rates (Bonenfant et al. 2009; Choquenot 1991; Van der
18 Wal et al. 2014). With rising density, individuals die from starvation because of
reduction in the availability and accessibility of food resources by conspecifics.
20 Agonistic interactions between individuals generate high stress levels which, in
turns, magnify the effect of other mortality sources (Peterson & Black 1988).
22 For instance, at high population density, epizootics are more likely to emerge and
to spread quickly in populations (). Higher densities may also force
24 non-dominant animals to occupy low-quality territories, ultimately decreasing
the average productivity and survival of the population (Krüger & Lindström
4
2001). Finally, because high population density exacerbates the effect of

2 environmental variation on demographic rates (Hones & Clutton-Brock 2007),
the risks of massive mortality events are much higher if the population is
4 maintained in this state on the long term. Given its numerous biological
consequences, an early detection of density-dependence is of the upmost
6 importance for the management and conservation of animal species to limit the
risk of catastrophic events.
8 The tamaraw (Bubalus mindorensis) is a large mammalian herbivore endemic
to the Island of Mindoro (Heude 1888; Custodio et al. 1996). Despite the
10 tamaraw being officially protected by law in 1954 and the creation of the Mts
Iglit-Baco Natural Park (MIBNP) protected area in 1970 (Maala 2001), its
12 distribution range has been dramatically shrinking over the last decades.
Originally found across the whole island of Mindoro with a gross estimate of
14 10,000 individuals in 1900 (Long et al. 2018), the tamaraw has suffered from
intensive land conversion for agriculture and logging industry, trophy hunting,
16 disease outbreaks spread out by domestic cattle (Maala 2001), and traditional
hunting conducted by upland indigenous communities who share their living
18 space with the large herbivore (Long et al. 2018). As a consequence, the species
is now found in a few isolated populations scattered across Mindoro, with
20 MIBNP hosting the largest number (Matsubayashi et al. 2010; Wilson &
Mittermeier 2011; Long et al. 2018). The MIBNP tamaraw population has been
22 restricted to one location of less than <3,000 ha in 2018, that we referred to as
the Core Zone of the Monitoring (CZM, see Fig. 1).
24 Given its critical importance for the conservation of this iconic species, the
MIBNP tamaraw population has been the cornerstone of all past, present and
5
future conservation plans (Maala 2001; Long et al. 2018). A better knowledge
2 about its ecology and population dynamics is fundamental to assess its current
conservation status, to strengthen protection measures and to guide future
4 conservation strategy with evidence-based information (Caughley 1994; Norris
2004; Stewart et al. 2005; Sutherland et al. 2004). Unfortunately, the population
6 dynamics of tamaraws in MIBNP has not been documented so far, nor do we
have any estimation of its average population growth rate. Yet it is likely that
8 density-dependence of demographic parameters will affect the population in the
near future because hunting and human disturbance prevent this population from
10 expanding outside of the CZM.
Here we took advantage of a 19 year-long time series of abundance data to
12 present the first detailed analysis of the tamaraw population dynamics in the
CZM of the MIBNP. We test the hypothesis that this population of large
14 herbivores could be facing negative effects of density. We predict (i) that because
of the substantial conservation effort in the last decade, overall population
16 growth rate (noted r) should be statistically > 0. Yet (ii), because the no-hunting
agreement area creates a spatial demarcation between risky and safe habitats, we
18 expect local growth rates to differ in space and predict a decrease of r while
getting away from the main rangers’ base camps and patrolling routes. Finally
20 (iii), given the absence of direct threats, we predict that the MIPBN tamaraw
population could be approaching the carrying capacity of the CZM and expect a
22 negative association between r and tamaraw abundance within the NHZ.
6
MATERIAL AND METHODS
2 Study site
The study area is located within the Mounts Iglit-Baco Natural Park (N12◦ 540
4 E121◦ 130 ) in the south-central part of Mindoro which, with 106,665 ha, is the
island largest protected area. MIBNP shelters the largest number of tamaraw in a
6 single site located on the south-west edge of the protected area (Fig. 1).
Tamaraw distribution range is now restricted to less than 3,000 ha, which
8 represents approx. 3% the MIBNP surface. This area is the Core Zone of the
Monitoring because this is where most of the patrolling and observations are
10 carried out by the Park’s officers. There are currently 15 park rangers divided
into two or three teams to monitor the area on a regular basis, following regular
12 distinct patrolling routes centered around the three base camps.
In an attempt to reinforce tamaraw protection, wildlife managers came to a
14 much needed agreement with residing indigenous people in 2016, declaring a
1,600 ha no-hunting zone area (NHZ) within the CZM (Fig. 1). An unforeseen
16 consequence of the NHZ establishment for tamaraw seems to be an increase of
pressure by traditional hunting at its direct boundaries. MIBNP rangers regularly
18 report setting of snare and spear traps, and violation of the NHZ agreement in
the last year is not uncommon with several deaths of tamaraws confirmed
20 (Boyles comm. pers.). Another possible response of large herbivores to hunting
is space use modification to reduce the risk of being killed (Chassagneux et al.
22 2019; Marantz et al. 2016).
The CZM area is a rolling grassland plateau with an average elevation of
24 800m a.s.l., dominated by cogon (Imperata cylindrica) and wild sugarcane
7
(Saccharum spontaneum), and interspersed with numerous wooded creeks,

2 secondary forest fragments and steep hills. In recent years, the invasive Siam
weed (Chromolaena odorata, locally called hagonoy) has been quickly
4 spreading in the grassland and now covers an unknown fraction of the tamaraw
habitat. The tropical climate is strongly seasonal with a rainy season spanning
6 from June to October (mean temperature 26.5◦ C; mean rainfall: 150 mm), and a
hot dry season from November to May (mean temperature 29.8◦ C; mean rainfall:
8 5 mm). Apart from humans, the only potential predator to the tamaraw is the
reticulated python (Malayopython reticulatus) that is capable of preying upon
10 calves or yearlings but no such observation have been made so far.
Tamaraw counts
12 Every year since 2000, the Tamaraw Conservation Program (TCP), a high profile
program of the Philippine Department of Environment and Natural Resources
14 (DENR), has been conducting tamaraw counts across the CZM in MIBNP,
yielding an annual index of relative population abundance. The tamaraw
16 counting area consists in 18 vantage points, covering nearly 2,200 ha within the
CZM (Fig. 1). The method is rather invasive for the MIBNP ecosystem because
18 it involves the burning of grasslands a few weeks ahead of each count, in order to
increase visibility and to attract tamaraws to areas with highly palatable and
20 attractive regrowing grasses. Each year, between 1,200 and 1,500 ha of
grasslands were intentionally burned for the purpose of monitoring tamaraw
22 abundance.
Individual tamaraws were counted simultaneously at the 18 vantage points
24 for two hours at dusk and dawn, and for four consecutive days, in late March or
8
early April. For the 8 sessions and at each vantage points, at least two observers
2 spotted and recorded all animal seen on field paper sheets, writing down
observation time, approximate location, and assessed sex and age-category of
4 individuals (split into calves, yearlings, sub-adults and adults). Right after the
completion of the 8 sessions, count data were cross-checked and cleaned for
6 obvious double observations, and the total number of different animals seen was
then derived for each vantage point s and year t. These annual counts make the
8 only available source of information about tamaraw population trends and
dynamics for officials and conservation agencies.
10 Data analyses
For populations of long-lived and endangered species, robust and accurate

12 estimations of r remains difficult to obtain. Knowledge about r requires either
the estimation of demographic rates combined into a Leslie matrix (Leslie 1945)
14 or the availability of long time series of population abundance, preferably on an
annual basis. In the absence of natural marking, capturing individuals of an
16 endangered species to make them identifiable with tags often face strong and
justified resistance for ethical reasons, mostly because of unavoidable mortality
18 risks. Consequently, in the best case the monitoring of the population abundance
of endangered species relies on an estimator accounting for detection probability
20 of animals (e.g. Kéry & Schmidt 2008), but most of time a simple index of
abundance is carried out.
22 Working with abundance index such as annual counts to derive an empirical
estimates of r however comes with some serious methodological issues (Îto
24 1972; Lindley 2003). Over the last decades, population counts have repeatedly
9
been associated with a large sampling variance (e.g. Caughley 1977), and
2 coefficient of variation for population abundance as high as 30% were reported
for species 1, species 2 for instance. From a conservation biologist viewpoint, a
4 large sampling variance in the input data for estimating r is a major pitfall
because of the over-optimistic results it leads to. Both the point estimate and its
6 precision are overestimated in the presence of sampling variance (Lindley 2003).
Despite the availability of appropriate statistical methods to account for the
8 sampling variance of population counts (Kalman filter, De Valpine & Hastings
2002), such tools remain seldom used in practice for conservation.
10 Here we analyzed the number of observed tamaraws and judged as different
individuals by observers for each of the 18 vantage points, split by years (Cs,t ,
12 where indice s ∈ {1, . . . , 18} stands for the vantage point, and
t ∈ {2000, . . . , 2019} stands for the time in years. We implemented a state-space
14 model to tease apart process from sampling variances, making the assumption
that double and under-counts compensate and are randomly distributed from
16 year to years. We worked in a Bayesian framework (e.g. Kéry & Schaub 2011)
but note this is fully equivalent to a Kalman filter with a frequentist approach. To
18 do so we defined Ns,t as the unobserved ’true’ abundance linked to Cs,t by a
random effect corresponding to the process variance (σc2 ), on the log scale to
20 ensure positive values for abundance. Our baseline model was:
log(Cs,t ) ∼ N (log(Ns,t ), σc2 )
Accordingly, we computed the population growth rate rs,t of the tamaraw

22 population at each vantage points s, as:
10

Ns,t+1
rs,t = log ,
Ns,t
from counts corrected for the sampling variance, hence returning an unbiased
2 estimation of the population growth rate. The overall population growth of the
tamaraw population of MIBNP is then given by:
1 19

Nt+1
r̄ = ∑ log Nt ,
19 t=1
4 whereby Nt = ∑18
s=1 Ns,t . We informed Nt with the total number of animals seen
each count session (Ctot ) and linked counts at the VP and population levels with
6 the following observation equation: Ctot ∼ N (Nt /p, σtot
2 ). The consolidated
number of tamaraws Ct being much smaller than total number seen Ct ot from
8 which double-counts were removed, we included a constant proportion
parameter p, similar to a ”detection rate” (which is not). By doing so, we could
10 use the available information about tamaraw counts fully, including both the
knowledge of experienced rangers and the raw number of detected tamaraws
12 with no a priori interpretation. In addition, we could input missing values at the
VP level for years 2000 to 2003 and 2005 thanks to the Ctot constrain on Nt .
14 To test our third hypothesis we investigated potential density-dependence in
the annual growth rate by fitting a logistic growth model to the total number of
16 tamaraws seen per year Nt :

Nt
log(Nt+1 ) = log(Nt ) + rm × 1 − ,
Kl
11
where rm is the maximum population growth for the tamaraw, and K is the
2 carrying capacity of the MIBNP. Likewise, we tested if the annual population
growth rate would decrease in time with population density by fitting a
4 Gompertz model:

KG
log(Nt+1 ) = log(N1 ) + log × 1 − e−β ×t ,
N1
Because the pair of parameters rm and Kl (logistic), and β and KG

6 (Gompertz), are not fully separately identifiable (Lebreton & Gimenez 2013), we
used informative priors for rm and β based on the reported maximum population
8 growth for other large bovids (see Traill et al. 2007, for a similar approach).
Accordingly, we set a moderately informative prior value of mean 0.3 and a
10 precision of 0.1 in our models (Table 2). We assessed the different models by
computing explicitly, during the model fit, the probability for our abundance
12 time series to be generated by each of the three demographic models
(exponential, Gompertz and logistic).
We fitted our statistical model to the tamaraw count data with JAGS 4.0
(Plummer et al. 2003), with 3 MCMC chains and a burn-in of 40,000 iterations.
We obtained parameter estimates with an additional 30,000 iterations and a
thinning factor of 5 (hence a total of 6,000 MCMC samples). We checked model
convergence graphically by investigating the mixing of MCMC chains, along
with the R̂ statistics (Brooks & Gelman 1998). With the exception of rm (see
above), we set uninformative priors with large variance for all other parameters
to be estimated, and checked the sensitivity of our results to the average value of
the priors by replicating the analyses with variable mean and initial values for
12
prior distributions. We report all parameters estimates as the mean of the

posterior distributions along with the 95% percentile for the credible intervals
following Louis & Zeger (2008): 95%low estimate 95%up .
Note that despite the fact that we analysed the number of seen tamaraws as
2 described above, we chose not to display such numbers on figures and to report a
standardized abundance (Cz,t /max(Cz,t )) instead. This transformation applies to
4 the estimated carrying capacities of the Gompertz and logistic models,
K/max(Cz,t ), hence reported values for K are > 1. We did not provide raw
6 counts to avoid a likely confusion between the index of relative tamaraw
abundance we worked with, and a real population size estimator that was not
8 implemented at MIBNP. Such a variable transformation has no incidence on the
biological interpretation of the results though.
10 RESULTS
From our baseline model of tamaraw counts, we estimated an average annual

12 growth rate of r̄ = 0.02 0.06 0.12 for the MIBNP population from 2000 to 2019,
which associated variance process equals σ proc = 0.25 0.37 0.49 (Fig. 2). As
14 expected for count data, the sampling variance was large σc = 7.32 9.31 9.98
suggesting frequent double counts of the same individuals. That the MIBNP
16 tamaraw population increased in size by c.a. 6% per year and was statistically
different from 0 is a clear support of our first hypothesis predicting increase of
18 abundance thanks to successful protection efforts at MIBNP.
Considering the vantage point specific growth rates (rs ), we observed a
20 marked variability in space (Table 2; Fig. 3) considering that growth rates ranged
13
between rs = −2.52 −0.32 0.20 at Tarzan VP and rs = 0.03 0.19 0.35 at Inubon VP.
2 Vantage point specific growth rates were spatially structured at MIBNP (Fig 3).
With no exception, rs < 0 were located at the periphery of the NHZ (Iyan,
4 Tarzan, Saligue East, Mibluan, Fangandatan) although rs of a few vantage points
was > 0 despite being located on the NHZ limit too (Nagbobong, Malibayong,
6 Talafu West and East). Together these results are in accordance with our
hypothesis of an on-going contraction of the tamaraw range at MIBNP after
8 2000, and the disappearance of individuals at the periphery of the CZM.
Analytical evidence for density-dependence in the tamaraw dynamics
10 between year 2008 and 2019 was moderate to high. We explicitly estimated the
probability π that the exponential, Gompertz and logistic models would be the
12 underlying process generating the observed variation in tamaraw abundance
(proxy of Baye’s factor). Our different models did have different probability to
14 have generated to observed data with, respectively π = 0.80, π = 0.00 and
π = 0.20 for the exponential, Gompertz and logistic model respectively. The
16 estimated coefficients of the Gompertz model lent statistical support for the
hypothesis of a gradual decrease in the annual population growth with time
18 (β = 0.07 0.09 0.11 ), and estimated carrying capacity to KG = 1.12 1.33 1.64 .
Similarly, point estimates for the two parameters of the logistic growth function
20 were rm = 0.09 0.11 0.13 for the maximum population growth rate and
Kl = 1.14 1.35 1.61 for the carrying capacity, very close to KG estimates.
14
DISCUSSION
2 The isolated tamaraw population in MIBNP has been increasing in size since
2000 (Fig. 2). Undoubtedly, for such an highly endangered species a positive
4 growth rate of 0.06 over 19 years of monitoring is a great success, both locally
and for the global conservation effort. Yet, the devil lies in the details though,
6 and two additional results darken this otherwise positive picture of tamaraw
population dynamics and conservation. First, our results highlight a progressive
8 decrease of the average growth rate, suggesting density-dependent effects at the
population level. Furthermore, we found support for the hypothesis of a
10 contraction of the tamaraw distribution, evidenced by a clear spatially structured
pattern of the population growth rates. The population dynamics we document is
12 clearly calling for new and urgent conservation actions if the future of tamaraw
is to be ensured.
14 Long-run Positive Growth of Tamaraws
The average population growth rate of r̄ = 0.06 we found gives strong evidence
16 in favor of our hypothesis of successful conservation policies of tamaraw over
the past two decades. Yet, an annual growth of 6% is relatively low compared to
18 other similar-sized or even larger cattle (Table 2). For instance, a population of
African buffalo (Syncerus caffer) in Southern Africa grew at a rate of 12% per
20 year for 28 years (Jolles 2007), and values close to 30% per year have regularly
been recorded for bison (Bison bison) or banteng (Bos javanicus) despite these
22 species are between 2 and 2.5 times larger in size than the tamaraw. From this
brief comparative approach, we could expect a substantially higher average
15
growth rate for tamaraws if the population was in a colonizing phase. Given the
2 tamaraw is threatened, our results suggest that specific environmental factors are
limiting its dynamic in MIBNP.
4 A first limiting factor could stand in the fact that the most available and
accessible biotope for the tamaraw in MIBNP is grassland (83% of the CMZ)
6 dominated by strong pioneer competitive species such as cogon and wild
sugarcane; both grasses becoming de facto prominent in the tamaraw diet. While
8 providing tamaraws with abundant and high quality forage at the early stages of
plant growth, the grassland could rapidly become of much lower nutritive value
10 during the dry season. Once grass growth is complete, stems exceeds tamaraw
height (300cm vs. < 120 cm respectively) and dry out to become unpalatable,
12 raising the question of food resource accessibility and availability for a large
population of tamaraws.
14 Dry vegetation also questions the ambivalent role of fire regime. On the one
hand, regular burning rejuvenate vegetation and promote regeneration of
16 attractive new grass shoots few months ahead of the rainy season, when animals
might look for alternative marginal habitats such as forest or creeks for
18 complementary feeding purposes. On the other hand, previous observation
suggest that regular fire could promote Siam weed expansion (Nath et al. 2019),
20 which is indeed rapidly spreading on the study site. Like other grazing
herbivores (Rozen-Rechels et al. 2017), tamaraw avoid this invasive plant, an
22 issue that could ultimately decrease the CMZ carrying capacity (Dumalisile &
Somers 2017). More generally tamaraw feeding preference is not fully
24 understood and the species might be a mixed-feeding or a browsing herbivore
(sensu Hofmann 1989). While being de facto restricted on grassland habitats,
16
tamaraw are suspected to be ecologically flexible, as recent assessment of a new

2 tamaraw population in mountain forest habitats in the upland inner Mindoro
Island would suggest (Schütz 2019). In other words, the CZM might actually
4 provide ecological conditions whose intrinsic limits are artificially reduced, so
far, thanks to long lasting and highly disturbing habitat management
6 policies;detrimental impact starting progressively to exceed beneficial effects.
Possible Impacts of Human Disturbance and Hunting
8 An additional factor, related to human activities, could also account for the low
tamaraw population growth rate we report. Tamaraw has always been
10 traditionally hunted by IPs and despite agreement of mutual effort of
conservation, both intentional and unintentional casualties still happen. Besides
12 tamaraw is sought after for bushmeat by lowlander poachers. Both threats are
hard to control in the field, are likely pervasive across MIBNP, and lead to an
14 increased mortality of all age-classes of tamaraws to an unknown extent.
Positive growth over the past two decades is a clear sign of successful protection
16 measures thanks to the constant presence of rangers and the recurrent patrols that
have likely contributed to decrease harvesting pressure within the CZM.
18 However, the marked spatial heterogeneity we document on the observed growth
rates with negative values at the periphery of the counting area might reveal a
20 lower protection efficiency when moving away from the main patrolling routes
and base camps. Actually, the CZM area functions like a source-sink system
22 with central areas nearby base camps concentrating most of the vantage points
with positive growth rates, with local value as high as 16% (Table 1), more in
24 line with the average growth we review for large bovids (Table 2).
17
Space Needed for Sustaining Population Growth
2 At first glance, the gradual decrease of the tamaraw population growth rate over
the survey coupled with the overall abundance observed at MIBNP reflects the
4 signature of some sort of density-dependence: a change in demographic rates
lowering the number of individuals added to the population from year to year
6 (Fowler 1987; Sinclair 1989). Predicted growth rates from the lowest to the
highest abundance varied between rt = 0.11 and rt = 0.01 for tamaraw at
8 MIBNP. Detection of density-dependence in MIBNP is worrying because it is
usually the sign of increased competition between individuals for food or
10 territorial space. In other word, our results suggest that tamaraws could already
be suffering from food resource shortage. Similar to the tamaraw case is the
12 banteng (Bos javanicus) population of Alas Purwo National Park, Indonesia. In
spite of an increasing abundance, this population has become consolidated close
14 to an artificially maintained grazing area, which may be causing increased
predation pressure of juveniles, and the population dynamics and carrying
16 capacity are not yet known (JB, pers. obs).
In absence of safe corridors for individuals to safely emigrate from the CZM
18 and in the presence of adverse effects of Siam weed on forage quantity, we
cannot expect this population to keep on increasing in size for the next decade as
20 we report between 2000 and 2019. From current data, we could estimate a
carrying capacity of 1.4 meaning the current MIBNP tamaraw abundance has
22 already reached almost 2/3 of the carrying capacity. This explanation is the
biological hypothesis of density-dependence, which is well known and
24 repeatedly detected in many populations of large mammals (Bonenfant et al.
18
2009). Yet, in absence of trustful demographic rates estimations of tamaraws at

2 MIBNP we hit the limits of the so-called pattern-oriented approach (Krebs
2002), with no way to assert real changes in demographic rates with abundance.
4 Consequently, an alternative interpretation of the density-dependence pattern
that we report for tamaraw is possible, which we describe as the count bias
6 hypothesis. Visual counts of animals, as it is conducted in MIBNP, are
notoriously known to be inaccurate and prone to underestimation of the real
8 abundance (Andersen 1953; ?) (Andersen 1953; Laake et al. 1999). Tamaraw
counts were not designed to be used with a population abundance estimator such
10 as returned by the distance sampling (Buckland et al. 2004) or capture-recapture
methods (Schwarz & Seber 1999), and therefore failed to account for the
12 imperfect detection of individuals. The proportion of undetected tamaraws could
increase with their overall abundance, a previously reported effect on red deer
14 (Cervus elaphus Garel et al. 2010) and roe deer (Capreolus capreolus, Pellerin et
al. 2018). Alternatively, different cognitive biases (Baddeley et al. 2004) could
16 generate similar patterns if, for instance, observers unconsciously pull the total
number of animals downward when consolidating count data. At this stage,
18 counts at year t could likely influence the results of counts at year t + 1,
assuming observers are conservative in their expectation of the tamaraw annual
20 population growth. The monitoring of additional biological variables in the field
such as the reproductive success of females could help at teasing apart the two
22 hypotheses (Morellet et al. 2007).
19
Implication for tamaraw conservation
2 Current conservation measures assume tamaraw population growth will continue

in the next decade, without a need to mitigate increased pressures thought to be
4 impacting the periphery of the CZM or to address the density dependence
pressure on the population. Our results call for a adjustment to strategic planning
6 for protection and management within MIBNP, as part of the MIBNP
Management Plan and the island-wide Tamaraw Conservation Management and
8 Action Plan (DENR 2019). Firstly, two main aspects must be urgently
addressed: (i) increase the available space for the species so to mitigate
10 density-dependence effect and maintain positive growth rate, that must be
combined with (ii) increasing the effectiveness of ranger patrols in the periphery
12 area of the CZM to reduce the recurrent poaching pressure from Filipino
lowlanders and losses due to traditional hunting. Both targets request to
14 collaborate closely with residing IP communities on a more sustainable land-use
and traditional hunting system. Indeed, a more effective protection and suitable
16 habitat management might in turn increase the average growth rate at MINBP,
and exacerbate the observed density-dependence demographic responses of
18 tamaraws. Secondly, when investigating the feasibility of translocating animals
from this source population to other sites, estimated demographic rates should be
20 incorporated into population modelling for extraction number that maintain a
sustainable yield and the tamaraw population viability. Indeed, the maximum
22 sustainable yield is found at K/2 in density-dependent models, not when
population abundance reaches carrying capacity K.
20
Conclusion
2 Despite increased efforts of land conservation, range contractions and habitat

fragmentation are still one of the most important threat to large animals
4 population (Ripple et al. 2015). As a consequence, a greater proportion of wild
herbivores (and their natural predators) are expected to be impacted by
6 intensively managed landscapes. So far, conservation responses essentially
consist in protecting large areas of land, a more than praiseworthy approach, that
8 nevertheless have limits, as illustrated by our results. In any case, positive
growth rates detected for a species in a conservation area could potentially hide a
10 darker picture. First, the apparent growth of a population, trapped in a too small
conservation area, can be the pervasive effect of range contraction rather than the
12 effect of increased reproduction and survival. Second, because of this, the
population could actually face important demographic challenges that, in the
14 long term, could lead to new threats. In this context, there is an increasing need
to understand the interaction between anthropogenic activities and the population
16 dynamics of protected species. Clearly, the protection of large herbivores may
hence come at the costs of undesirable ecological effects, rendering their
18 conservation even more difficult. On the bright side, our findings also
exemplifies how detailed analyses of relative population abundance data, even
20 without absolute population size estimators, can contribute to determining future
conservation measures and are essential for large mammal conservation.
21
ACKNOWLEDGMENTS
2 A lot of people are to thank here. We are grateful to Fernando Garcı̀a for producing
maps used in this paper. Special thanks are extended to Alvaro Gonzalez. We must
4 acknowledge the invaluable help of all rangers, students and volunteers who contributed
to the annual tamaraw counts.
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TABLE CAPTIONS
2 Table 1 Annual growth rate (r ) of tamaraw abundance as estimated at

each vantage counts in Mounts Iglit-Baco Natural Park located on
4 Mindoro island, Philippines, between 2006 and 2019. We used a
Bayesian state-space model to account for sampling variance in animal
6 counts and reported the posterior mean estimation and its associated
95% credible interval (CI). Note the marked spatial variation in the
8 temporal trend of tamaraw abundance, ranging from a rapid decline at
Tarzan to a marked increase at Bato Fidel or Inubon (see also Fig 3).
10 Table 2 Estimated annual growth rates (r) of some large bovinae

populations with body size comparable to the endangered tamaraw
12 (Bubalus mindorensis). The observed population growth rate of
tamaraws at Mounts Iglit-Baco Natural Park, Mindoro, Philippines,
14 between 2000 and 2019 of r = 0.06 lies at the lower range of reported
values.
29
Table 1
Vantage point r 2.5% 97.5%

Loibfo 0.12 −0.01 0.25
Magawang 0.12 −0.02 0.24
Bayokbok 0.12 −0.01 0.25
Bato Fidel 0.15 0.01 0.27
Inubon 0.15 0.02 0.28
Mibluan 0.13 −0.01 0.26
Nagbobong 0.02 −0.14 0.19
Fangandatan −0.03 −0.24 0.14
Anyayos 0.08 −0.06 0.21
Lanas I 0.10 −0.05 0.24
Lyan −0.02 −0.24 0.16
Tarzan −0.18 −0.42 0.02
Talafu East 0.06 −0.10 0.20
Talafu West 0.07 −0.08 0.21
Malitwang 0.08 −0.08 0.22
Lanas II 0.02 −0.13 0.17
Saligi East −0.18 −0.45 0.04
Malibayong 0.02 −0.16 0.19
Saligi East −0.46 −2.98 0.17
30
C. Bonenfant
Table 2
Species Location r Reference

Bison bison Mackenzie Bison Sanctuary, Canada 0.21 Gates & Larter (1990)
Bison bison Mackenzie Bison Sanctuary, Canada 0.20 Larter et al. (2000)
Bison bison Yellowstone, Wyoming, USA 0.07 Fuller et al. (2007)

Bison bison Yellowstone, Wyoming, USA 0.30 Singer & Norland (1994)
Bison bonasus Białowieża, Poland 0.07 Mysterud et al. (2007)1
Bison bonasus Białowieża, Poland 0.18 Krasiński (1978)1
Bos gaurus Thung Yai Naresuan Sanctuary, Thailand 0.31 Steinmetz et al. (2010)
Bos javanicus Not applicable, model prediction 0.32 Hone et al. (2010)
Bos javanicus Cobourg peninsula, Australia 0.00 Choquenot (1993)
Boselaphus tragocamelus Kanha National Park, India 0.18 Mathur (1991)
Boselaphus tragocamelus Keoladoe National Park, India 0.02 - 0.06 Haque (1990)
Bubalus arnee 0.02 - 0.06 Khatri et al. (2012)
Bubalus arnee Koshi Tappu Wildlife Reserve, Nepal 0.03 Heinen & Paudel (2015)
Bubalus bubalis 0.03 Heinen (1993)
Bubalus bubalis ? Freeland and Boulton (1990)
Bubalus mindorensis Mount Iglit-Baco National Park, Philipines 0.06 This study
Tamaraw dynamics on Mindoro

Pseudoryx nghetinhensis Vietnam & Laos 0.05 Kemp et al. (1997)b
Syncerus caffer Hluhluwe-iMfolozi Park, South Africa 0.12 Jolles (2007)
Syncerus caffer Zakouma National Park, Chad 0.08 Cornélis et al. (2014)
Taurotragus oryx Lombard Nature Reserve, South Africa 0.12 Buys & Dott (1991)c
Taurotragus oryx South Africa 0.16 Van Houtan et al. (2009)
Taurotragus oryx Kruger National Park, South Africa −0.02 Nicholls et al. (1996)
31
Taurotragus oryx Masai Mara Ecosystem, Kenya −0.11 - 0.07 Ottichilo et al. (2000)
Taurotragus derbianus Senegal 0.31 Koláčková et al. (2011)
FIGURE CAPTIONS
2 Figure 1 General location of the Mounts Iglit-Baco Natural Park on Mindoro
island, Philippines. The current geographical range of the endangered tamaraw
4 (Bubalus mindorensis) population is limited to a 2,500ha area (salmon color
area) of which 1,250ha consists in a no-hunting zone (red dotted line) since
6 2016, hence offering the highest level of protection for the species. Every year
since 2000, taramaw counts have been carried out in March–April on 18
8 different vantage points (green circles) for the purpose of monitoring population
abundance trends.
10 Figure 2 Time series of tamaraw (Bubalus mindorensis) abundance in Mounts
Iglit-Baco Natural Park on Mindoro island, Philippines, from years 2000 to 2019.
12 Raw counts (black solid line) and counts corrected for sampling variance (blue
solid line) are presented with the associated 95% credible interval. Over the 13
14 years of monitoring, tamaraw abundance increased at a rate of 6% per year on
average.
16 Figure 3 Spatial variation in the local growth rate of tamaraw (Bubalus
mindorensis) abundance in Mounts Iglit-Baco Natural Park on Mindoro island,
18 Philippines, from years 2000 to 2019. With a general positive growth since 2000,
the situation in the core area of the monitoring zone is in sharp contrast with the
20 important decrease in abundance observed at the periphery. Movement of
animal toward less disturbed areas or illegal hunting outside of the no-hunting
22 agreement zone could account for this marked spatial structure of the tamaraw
population dynamics.
32
Figure 1
121°3′0.000″ 121°4′48.000″ 121°6′36.000″

12°43′12.000″
L egend
Items
Vantage Points
Ranger Station
IP Sitio
Mountain
Management and
Monitoring Zones
12°41′24.000″
Annual Count Area

Agreed Free Hunting
Zone 2016 (1600 Ha)
2017 Maximum Tamaraw
Distribution (2500 ha)
33
Figure 2
1.0
●
●
●
Relative Tamaraw Adundance
0.8
●
●
●
●
● ●
●
●
0.6 ●●
●
●
●
● ●
●
Legend:
● ●
●
●
●
● ● ●
● Obs.
0.4 ● Pred.
2010 2015
Time (in years)
34
Figure 3
121°3′0.000″ 121°4′48.000″
12°43′12.000″
L egend
Items
Ranger Station
Mountain
Management and
Monitoring Zones
Annual Count Area
Agreed Free Hunting
Zone 2016 (1600 Ha)
Annual Growth Rate
12°41′24.000″
-0,24 ; -0,20
-0,20 ; -0,15
-0,15 ; -0,10
-0,10 ; -0,05
-0,05 ; 0,00
0,00 ; 0,05
0,05 ; 0,10
0,10 ; 0,12
35

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Cast away on Mindoro island: population

C. Bonenfant1 , A. Rutschmann2 , J. Burton3 , R. Boyles4 , and

C. Bonenfant Tamaraw dynamics on Mindoro

2 Endangered species, despite living at low population density, may undergo

C. Bonenfant Tamaraw dynamics on Mindoro

2 In conservation planning, population abundance is critical and focuses the

C. Bonenfant Tamaraw dynamics on Mindoro

C. Bonenfant Tamaraw dynamics on Mindoro

2001). Finally, because high population density exacerbates the effect of

C. Bonenfant Tamaraw dynamics on Mindoro

C. Bonenfant Tamaraw dynamics on Mindoro

MATERIAL AND METHODS

C. Bonenfant Tamaraw dynamics on Mindoro

(Saccharum spontaneum), and interspersed with numerous wooded creeks,

C. Bonenfant Tamaraw dynamics on Mindoro

For populations of long-lived and endangered species, robust and accurate

C. Bonenfant Tamaraw dynamics on Mindoro

log(Cs,t ) ∼ N (log(Ns,t ), σc2 )

Accordingly, we computed the population growth rate rs,t of the tamaraw

C. Bonenfant Tamaraw dynamics on Mindoro

C. Bonenfant Tamaraw dynamics on Mindoro

Because the pair of parameters rm and Kl (logistic), and β and KG

C. Bonenfant Tamaraw dynamics on Mindoro

prior distributions. We report all parameters estimates as the mean of the

From our baseline model of tamaraw counts, we estimated an average annual

C. Bonenfant Tamaraw dynamics on Mindoro

C. Bonenfant Tamaraw dynamics on Mindoro

14 Long-run Positive Growth of Tamaraws

C. Bonenfant Tamaraw dynamics on Mindoro

C. Bonenfant Tamaraw dynamics on Mindoro

tamaraw are suspected to be ecologically flexible, as recent assessment of a new

Possible Impacts of Human Disturbance and Hunting

C. Bonenfant Tamaraw dynamics on Mindoro

Space Needed for Sustaining Population Growth

C. Bonenfant Tamaraw dynamics on Mindoro

2009). Yet, in absence of trustful demographic rates estimations of tamaraws at

C. Bonenfant Tamaraw dynamics on Mindoro

Implication for tamaraw conservation

2 Current conservation measures assume tamaraw population growth will continue

C. Bonenfant Tamaraw dynamics on Mindoro

2 Despite increased efforts of land conservation, range contractions and habitat

C. Bonenfant Tamaraw dynamics on Mindoro

to the annual tamaraw counts.

22 Caughley, G. (1977). Analysis of vertebrate populations. Wiley, London. Graeme

Caughley, G. (1994). Directions in conservation biology. Journal of animal ecology, pp.

30 Choquenot, D. (1991). Density-dependent growth, body condition, and demography in

C. Bonenfant Tamaraw dynamics on Mindoro

Courchamp, F., Clutton-Brock, T. & Grenfell, B. (1999). Inverse density dependence

De Valpine, P. & Hastings, A. (2002). Fitting population models incorporating process

18 Eberhardt, L.L. (2002). A paradigm for population analysis of long-lived vertebrates.

20 Fowler, C.W. (1987). A review of density dependence in populations of large mammals.

C. Bonenfant Tamaraw dynamics on Mindoro

6 Hofmann, R.R. (1989). Evolutionnary steps of ecophysiological adaptation and

14 Îto (1972). On the methods for determining density-dependence by means of regression.

16 Jolles, A.E. (2007). Population biology of african buffalo (Syncerus caffer) at

Kéry, M. & Schaub, M. (2011). Bayesian population analysis using WinBUGS: a

Koláčková, K., Hejcmanova, P., Antonı́nová, M. & Brandl, P. (2011). Population